SCIENTIA MARINA 70 (3)
September 2006, 363-379, Barcelona (Spain)
ISSN: 0214-8358
Gelatinous zooplankton net-collected in the Gulf of
Maine and adjacent submarine canyons: new species,
new family (Jeanbouilloniidae), taxonomic
remarks and some parasites
FRANCESC PAGÈS 1, PER FLOOD 2 and MARSH YOUNGBLUTH 3
1
Institut de Ciències del Mar (CSIC), Passeig Marítim de la Barceloneta 37-49, 08003 Barcelona, Catalunya, Spain.
E-mail: [email protected]
2
Bathybiologica A/S, Bergen, Norway. E-mail: [email protected]
3
Harbor Branch Oceanographic Institution, Fort Pierce, Florida, USA. E-mail: [email protected].
SUMMARY: A systematic account is provided of the siphonophores, medusae, ctenophores, molluscs and salps net-collected in basins of the Gulf of Maine and adjacent canyons during three cruises carried out in September 2002, 2003 and
2004. We describe the siphonophore Lensia quadriculata sp. nov. and the hydroidomedusae Jeanbouillonia maserati gen.
nov. et sp. nov., the latter is the first member of the new family Jeanbouilloniidae. New observations are made about the morphology and/or systematics of the siphonophore Gilia reticulata and the hydroidomedusae Bougainvillia platygaster,
Bougainvillia principis, Modeeria rotunda, Clytia hemisphaerica, Orchistoma collapsum and Dichotomia cannoides.
Parasites associated with the ctenophores Pleurobrachia pileus (a cestode) and Beroe sp. (a hyperid amphipod) are reported.
Keywords: gelatinous zooplankton, Gulf of Maine, new species, Jeanbouillonia, Lensia, Gilia, Bougainvillia, Modeeria,
Dichotomia, Dipleurosoma.
RESUMEN: ZOOPLANCTON GELATINOSO RECOGIDO CON REDES EN EL GOLFO DE MAINE CON DESCRIPCIONES DE UN NUEVO SIFONÓFORO, UNA NUEVA ANTOMEDUSA Y DIAGNOSIS DE LA NUEVA FAMILIA JEANBOUILLONIIDAE. – Se presenta un directorio sistemático de los sifonóforos, medusas, ctenóforos, moluscos y salpas recogidos con redes en las cuencas del golfo de Maine y
cañones submarinos adyacentes durante tres cruceros realizados en septiembre de 2002, 2003 y 2004. El sifonóforo Lensia
quadriculata sp. nov. y la hidroidomedusa Jeanbouillonia maserati gen. nov. et sp. nov. son descritos, ésta última especie
es el primer miembro de la nueva familia Jeanbouilloniidae. Se aportan nuevas observaciones sobre la morfología y/o posición sistemática del sifonóforo Gilia reticulata y de las hidroidomedusas Bougainvillia platygaster, Bougainvillia principis,
Modeeria rotunda, Clytia hemisphaerica, Orchistoma collapsum y Dichotomia cannoides. Se registran algunos parásitos
asociados a los ctenóforos Beroe sp. (un anfípodo hipérido) y Pleurobrachia pileus (un céstodo).
Palabras clave: zooplancton gelatinoso, golfo de Maine, especies nuevas, Jeanbouillonia, Lensia, Gilia, Bougainvillia,
Modeeria, Dichotomia, Dipleurosoma.
INTRODUCTION
The epipelagic (0 to 200 m depth) and
mesopelagic (200 to 1000 m depth) gelatinous zooplankton living in the Gulf of Maine and adjacent
waters have mainly been researched in the second
half of the XIX century (e.g. Agassiz, 1865; Fewkes,
1888) and the first half of the XX century (e.g.
Mayer, 1912, Bigelow, 1914, 1926). Since then,
oceanographic research has shifted towards more
ecological studies, occasionally updating the gelatinous populations from in situ observations in sub-
364 • F. PAGÈS et al.
mersible dives (e.g. Larson et al., 1988). In
September 2002, 2003 and 2004, three cruises were
conducted in two basins (Wilkinson and Georges) of
the Gulf of Maine and four adjacent submarine
canyons (Oceanographer, Hydrographer, Atlantis
and Lydonia) (north-western Atlantic) (Fig. 1) in
order to explore the pelagic habitat and trophodynamics of the physonect siphonophore Nanomia
cara using the manned submersibles Johnson-SeaLink. Zooplankton samples were collected between
dives to complement the knowledge on the gelatinous populations living in the waters explored. The
present study reports on all the species net-collected
during the three cruises, describes two new cnidarian
species and provides observations on the morphology and/or systematics of other little-known species.
µm mesh-size) in surface waters. The Mocness
equipped with nine nets was hauled either horizontally or obliquely at selected depths. Most of specimens were sorted with the naked-eye after recovering the nets and they were immediately examined
and photographed under a binocular microscope or
while drifting in a Planktonkreisel. Afterwards the
samples were fixed in 4% buffered formaldehyde.
Marked differences in the sampling occurred
between cruises. For example, twenty Mocness
hauls were conducted from near the seafloor to the
sea surface among the three cruises, fifteen in 2002,
four in 2003 and only one in 2004 after the winch
broke. The six Meternet hauls were carried out in
2004. No attempt to make interannual variability
analyses was made due to this random sampling.
MATERIAL AND METHODS
RESULTS
The specimens were captured either by a
Meternet (200 µm mesh-size) hauled vertically and
a multi-net Mocness (300 µm mesh-size) in the
water column, and by a dipnet and a conical net (90
Eighty-one species of gelatinous zooplankton
were identified. In total, 33 were siphonophores, 28
hydroidomedusae, 8 scyphomedusae, 3 ctenophores,
6 molluscs and 3 salps (Table 1). One calycophoran
FIG. 1. – Map of the Gulf of Maine showing the basins and submarine canyons where zooplankton samples were collected.
SCI. MAR., 70(3), September 2006, 363-379. ISSN: 0214-8358
NEW SPECIES OF PLANKTONIC CNIDARIANS FROM THE GULF OF MAINE • 365
TABLE 1. – List of gelatinous species net-collected in the Gulf of Maine over the three cruises including those not mentioned in the main text.
Taxa (species number) and species name
CNIDARIA
SIPHONOPHORAE (33)
Agalma elegans
Agalma okeni
Nanomia cara
Cordagalma ordinata
Physophora hydrostatica
Amphycarion acaule
Praya reticulata
Rosacea flaccida
Desmophyes annectens
Hippopodius hippopus
Vogtia glabra
Vogtia spinosa
Sulculeolaria quadrivalvis
Chelophyes appendiculata
Diphyes bojani
Diphyes dispar
Dimophyes arctica
Lensia asymmetrica
Lensia conoidea
Lensia meteori
Lensia multicristata
Lensia quadriculata sp. nov.
Gilia reticulata
Crystallophyes amygdalina
Clausophyes ovata
Chuniphyes moserae
Chuniphyes multidentata
Abylopsis eschscholtzi
Abylopsis tetragona
Enneagonum hyalinum
Ceratocymba leuckarti
Ceratocymba sagittata
Sphaeronectes gracilis
ANTHOMEDUSAE (7)
Bougainvillia principis
Bougainvillia platygaster
Leuckartiara octona
Leuckartiara sp.
Cytaeis spp
Chromatonema rubrum
Jeanbouillonia maserati gen. et sp. nov.
LEPTOMEDUSAE (9)
Modeeria rotunda
Laodicea undulata
Clytia hemisphaerica
Clytia mccradyi
Zygocanna vagans
Aequorea sp.
Orchistoma collapsum
Dichotomia cannoides
Melicertum octocostatum
Location
Depth in metres (number of specimens)
Date
Georges Basin
Georges Basin
Oceanographer
Oceanographer
Atlantis
Georges Basin
Wilkinson Basin
Oceanographer
Oceanographer
Oceanographer
Atlantis
Oceanographer
Hydrographer
Oceanographer
Georges Basin
Oceanographer
Atlantis
Oceanographer
Oceanographer
Atlantis
Atlantis
Oceanographer
Georges Basin
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Hydrographer
Oceanographer
Oceanographer
Atlantis
Hydrographer
Atlantis
Hydrographer
10-0 (1)
35-7 (1)
350-0 (3)
500-375 (1)
600-0 (1)
280-0 (1)
75-0 (2)
surface (1)
225-150 (1)
375-2 (2)
600-0 (1)
35-0 (1)
172-80 (1)
500-375 (1)
surface (1)
350-0
600-0 (1)
350-0 (1)
150-6 (1)
501-464 (1)
600-0 (1)
35-0 (1)
surface (1)
677-629 (1)
surface waters
surface waters
350-0 (2)
35-0 (1)
678-578 (1)
777-678 (1)
500-375 (1)
100-0 (2)
678-578 (1)
100-0 (2)
678-578 (2)
10/9/03
11/9/03
16/9/04
23/9/03
21/9/04
25/9/04
27/9/04
15/9/04
2/9/02
25/9/03
21/9/04
16/9/04
13/9/02
23/9/03
13/9/03
16/9/04
21/9/04
16/9/04
26/9/04
16/9/02
21/9/04
16/9/04
24/9/04
9/9/02
15/9/04
15/9/04
16/9/04
16/9/04
13/9/02
9/9/02
23/9/03
20/9/04
13/9/02
20/9/04
13/9/02
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Hydrographer
Oceanographer
Oceanographer
Oceanographer
Hydrographer
Oceanographer
Oceanographer
Atlantis
777-678 (1)
777-678 (1)
777-678 (1)
661-600 (1)
678-578 (1)
35-0 (1)
35-0 (2)
350-0 (1)
317-278 (1)
35-0 (1 eudoxid)
0-661 (1 eudoxid)
100-0 (2)
9/9/02
9/9/02
9/9/02
16/9/04
13/9/02
16/9/04
16/9/04
16/9/04
17/9/02
16/9/04
16/9/04
20/9/04
Georges Basin
Wilkinson Basin
Oceanographer
Oceanographer
Oceanographer
Oceanographer
on surface (3)
35-2 (1)
200-100 (1)
350-0 (1)
777-728 (1)
750-625 (1), 500-375 (1)
13/9/03
10/9/03
16/9/04
16/9/04
9/9/02
15/9/03
Oceanographer
350-0 (1)
16/9/04
Atlantis
Oceanographer
Oceanographer
100-0 (2)
150-6 (2)
200-100 (1), 100-5 (3), 350-0 (1)
20/9/04
26/9/03
16/9/04
off Gloucester
55-0 (2)
9/9/04
Remarks
see text
see text
see text
see text
Fig. 7F
Fig. 7G
see text
see text
see text
see text
see text
SCI. MAR., 70(3), September 2006, 363-379. ISSN: 0214-8358
366 • F. PAGÈS et al.
TABLE 1 (Cont.). – List of gelatinous species net-collected in the Gulf of Maine over the three cruises including those not commented in the
main text.
Taxa (species number) and species name
LIMNOMEDUSAE (1)
Proboscidactyla mutabilis
TRACHYMEDUSAE (7)
Liriope tetraphylla
Rhopalonema velatum
Halicreas minimum
Aglantha digitale
Crossota rufobrunnea
Pantachogon haeckeli
Arctapodema ampla
NARCOMEDUSAE (4)
Aeginura grimaldii
Cunina fowleri
Cunina peregrina
Solmissus incisa
SCYPHOMEDUSAE (8)
Nausithoe atlantica
Nausithoe punctata
Atolla vanhoeffeni
Atolla wyvillei
Periphylla periphylla
Phacellophora camstchatica
Pelagia noctiluca
Poralia sp.
CTENOPHORA (3)
Tortugas Red Cydippid
(sensu G.R. Harbison)
Pleurobrachia pileus
Beroe sp.
MOLLUSCA (6)
Corolla calceola
Carinaria sp
Pterotrachea coronata
Pterotrachea hippopodius
Firoloida desmaresti
Phyllirhoe bucephala
TUNICATA (3)
Salpa fusiformis
Cyclosalpa affinis
Thalia democratica
Location
Depth in metres (number of specimens)
Date
Atlantis
100-0 m (2)
20/9/04
Oceanographer
Atlantis
Hydrographer
Oceanographer
Oceanographer
Oceanographer
Wilkinson Basin
Oceanographer
Hydrographer
Oceanographer
Oceanographer
Oceanographer
Oceanographer
350-0 (1)
501-464 (1)
370-278 (1)
750-625 (1)
500-375 (1)
998-875 (1)
75-0 (1)
998-875 (3), 875-672 (6)
878-777 (3), 777-680 (4), 680-576 (3)
998-875 (2), 875-672 (1), 625-500 (1)
661-600 (1)
600-0 (1)
900-800 (3)
16/9/04
16/9/02
17/9/02
15/9/03
15/9/03
23/9/03
27/9/04
23/9/03
17/9/02
23/9/03
16/9/04
16/9/04
3/9/02
Oceanographer
Hydrographer
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Atlantis
686-586 (3)
878-777 (1), 777-680 (1), 680-576 (1)
779-683 (8)
875-672 (3), 672-625 (3)
1000-625 (11)
350-0 (1)
surface waters (2)
683-779 (1)
750-625 (1), 625-500 (2)
684-636 (4)
998-875 (1)
750-625 (1)
600-0 (1)
5/9/02
17/9/02
8/9/02
23/9/03
25/9/03
16/9/04
15/9/04
8/9/02
15/9/03
22/9/02
23/9/03
25/9/03
21/9/04
Hydrographer
Oceanographer
Oceanographer
Atlantis
Oceanographer
Oceanographer
Oceanographer
Atlantis Canyon
Oceanographer
Oceanographer
Oceanographer
Oceanographer
Lydonia
Georges Basin
Atlantis
Atlantis
Hydrographer
878-777 (2)
625-500 (1)
998-875 (1)
124-88 (2)
350-0 (3)
875-750 (2), 750-625 (1)
600-500 (1)
600-0 (1)
625-500 (1)
778-680 (1)
850-750 (1)
600-0 (1)
surface waters (2)
210-185 (1)
100-0 (2)
600-0 (1)
878-777 (1)
17/9/02
15/9/03
23/9/03
16/9/02
16/9/04
25/9/03
16/9/04
21/9/04
15/9/04
5/9/02
25/9/04
16/9/04
22/9/03
20/9/03
20/9/04
21/9/04
17/9/02
Hydrographer
Oceanographer
Oceanographer
Oceanographer
700-600 (1)
681-631 (1)
625-500 (1)
998-875 (1)
17/9/02
22/9/02
15/9/03
23/9/03
Oceanographer
Oceanographer
Atlantis
Atlantis
Oceanographer
Oceanographer
375-2 (2)
100-5 (1)
124-88 (1)
124-88 (1)
surface waters (1)
100-5 (3)
25/9/03
Oceanographer
Atlantis
Nantucket Sound
Georges Basin
Georges Basin
Georges Basin
350-0 (many)
600-0 (many)
Surface (1 colonial)
35-0 (many), 60-35 (few)
85-60 (few), 110-85 (very few)
10-0 (many)
16/9/04
21/9/04
7/9/02
11/9/03
11/9/03
24/9/04
SCI. MAR., 70(3), September 2006, 363-379. ISSN: 0214-8358
16/9/02
16/9/02
15/9/04
16/9/04
Remarks
Fig. 7 H
Fig. 7 I
see text
see text
NEW SPECIES OF PLANKTONIC CNIDARIANS FROM THE GULF OF MAINE • 367
FIG. 2. – Anterior nectophore of Gilia reticulata, 5 mm high: A, lateral view. B, dorsal view. C, apical view. D, ostial view.
siphonophore and one anthomedusan were identified as new species. Both are described in the following systematic account that includes morphological descriptions of little-known species.
Family DIPHYIDAE Quoy and Gaimard, 1827
Subfamily GILIINAE Pugh and Pagès, 1995
Gilia reticulata (Totton, 1954)
(Fig. 2A-D)
Phylum CNIDARIA Verrill, 1865
Class HYDROIDOMEDUSAE Claus, 1877
Subclass SIPHONOPHORAE Eschscholtz, 1829
Order CALYCOPHORA Leuckart, 1854
Material: One anterior nectophore collected in Oceanographer
Canyon (40°11.8’N, 67°57.8’W) between 686 and 586 m depth on
5 September 2002. One anterior nectophore and three bracts collected in Oceanographer Canyon (39°52.7’N, 68°48.0’W) between
777 and 678 m depth on 9 September 2002. One anterior nectophore collected in Lydonia Canyon (40°19.8’N, 67°40.9’W)
between 678 and 634 m on 21 September 2002.
SCI. MAR., 70(3), September 2006, 363-379. ISSN: 0214-8358
368 • F. PAGÈS et al.
9 September 2002; seawater temperature 4.5 to 4.8°C, seawater
salinity 34.95 to 34.97 psu.
Additional material: One anterior nectophore collected by a
Mocness haul in Oceanographer Canyon waters (40°44.3’N,
68°02.6’W) in the 998 to 875 m depth range on 23 September 2003.
Three anterior nectophores collected at 1049 m depth in Suruga
Bay, Japan (34°47.1’N, 138°36.6’E) on 28 March 1985 in the horizontal haul 85131 during the cruise TO-84-23 conducted by the
Tokai University. One anterior nectophore collected at 790 m depth
in Suruga Bay on 13 November 1992 in the horizontal haul 92006
of the cruise TO 92-21. Holotype deposited at the Smithsonian
Institution. Paratypes deposited at the Smithsonian Institution,
Tokai University and the Institut de Ciències del Mar.
Etimology: quadriculata, small square, Latin,
feminine; from quadrus, square and culata, small.
FIG. 3. – Lateral view, slightly ventral, of the anterior nectophore of
Lensia quadriculata sp. nov. Nectophore height 6 mm.
Remarks: Pugh and Pagès (1995) established the
new genus Gilia when they described the eudoxid
stage of this reticulated species and wrote a complete description of the nectophores with detailed
drawings. The photographs of the anterior nectophore (Fig. 2) complement the previous study as
they show some little-known features of a pristine
specimen. The network of ridges forms a pattern of
hexagons and pentagons. Below the velar ridge, all
ridges are orientated in the vertical plane (Fig. 2AB) and the hydroecium is large and deep (Fig. 2D).
The nectophore perimeter (Fig. 2D) does not show
concave facets between the ridges, although the
ridge walls can cause this impression. The nectophore apex is rounded, not pointed. Cairns et al.
(2002) reported on this species only in the Pacific
coast of USA and Canada.
Subfamily DIPHYINAE
Lensia quadriculata sp. nov.
(Figs. 3, 4A-D, 5A)
Type material: Three anterior nectophores (one holotype and two
paratypes) collected by a Mocness haul in Oceanographer Canyon
waters (39°52.7’N, 68°48.0’W) in the 777 to 678 m depth range on
SCI. MAR., 70(3), September 2006, 363-379. ISSN: 0214-8358
Description: Anterior nectophore small and
very fragile, 6 mm high, 3 mm in maximum (central) dorso-ventral depth, 2.9 mm in maximum
(central) left-right width. The surface consists of
thin vertical and horizontal ridges crossing at right
angles and at equal intervals forming a quadrilled
pattern. Five sets (1 dorsal, 2 dorso-lateral and 2
ventro-lateral) of 3 longitudinal ridges each extend
from the pointed apex down towards the ostium,
which is reached only by the central dorsal ridge
and the dorso-lateral ridges. More longitudinal
ridges (2 ventral, 5 left ventro-lateral, 5 right-ventro-lateral, 3 left dorso-lateral and 3 right dorso-lateral) run from below the apex downwards, all
reaching the ostium except the 2 ventral ones. A
furrow grooves from the apex to the ostium
between the ventro-lateral and the dorso-lateral
ridges. Twenty ridges join the ostium. About 30
transversal ridges cross the longitudinal ones perpendicularly between the somatocyst level and the
apex giving rise to a characteristic quadrilled pattern of ridges. A constriction surrounds the nectophore at the somatocyst level but does not form a
velar ridge. The nectosac occupies the subumbrellar cavity almost completely. The somatocyst is
short, oval in shape, wider (1 mm) than it is high,
with a short pedicular canal, completely orange in
colour (Figs. 3, 4, 5A). The hydroecium is deep,
bell-shape, opened ventrally, with the apical surface above the ostium level. The mouth-plate is
composed of two basal lamellae slightly overlapped in the centre. The velum is very narrow.
A posterior nectophore partially distorted but
attributable to this species was also found in the
same sample that contained the holotype. However,
we are waiting for new material in good conditions
to be captured to make a correct description. The
eudoxid is unknown.
NEW SPECIES OF PLANKTONIC CNIDARIANS FROM THE GULF OF MAINE • 369
FIG. 4. – Holotype of Lensia quadriculata sp. nov., nectophore 6 mm high: A, ventro-lateral view. B, ventro-ostial view. C, detail of the
quadrilled pattern of crossed ridges near the apex. D, ostio-lateral view.
Remarks: Some years before these nectophores
were collected in the Gulf of Maine, the University
of Tokai in Shimizu (Japan) loaned FP a large collection of siphonophores sampled in the Bay of
Suruga over the period 1981 to 1992. Several new
species were identified in the collection and one of
them was the Lensia species described here. These
specimens were laterally flattened and it was decided to postpone the description until pristine ones
were captured.
The quadrilled pattern of crossed ridges suggests
that the new species could be phylogenetically clos-
er to Gilia reticulata (Fig. 2, see also Pugh and
Pagès, 1995). However, the multistriate diphyinae
Lensia ajax Totton, 1941, and L. lelouveteau Totton,
1941 are also quite close since both show the pattern
of five groups of longitudinal ridges. The marked
honeycomb arrangement of the ridges in G. reticulata is easily distinguishable from the cobweb pattern (radiating strand more prominent than the cross
connecting ones) of L. quadriculata. The distinction
is more difficult when comparing the nectophore of
L. quadriculata with preserved samples of L. ajax
and L. lelouveteau. Nevertheless, L. ajax somatocyst
SCI. MAR., 70(3), September 2006, 363-379. ISSN: 0214-8358
370 • F. PAGÈS et al.
FIG. 5. – A: Lensia quadriculata sp. nov., holotype in lateral view, nectophore 6 mm high. Bougainvillia principis, umbrella height 12 mm:
B, interradial view. C, perradial view. D, apico-lateral view. E, detail of the umbrella margin showing the “epaulettes” and the marginal bulbs
and tentacles. Bougainvillia platygaster, umbrella 5 mm high: F, perradial view. Jeanbouillonia maserati, gen. nov. et sp. nov., umbrella 3
mm high: G and I, lateral view; note a radiolarian on the apical exumbrellar surface. H, trident-shape distal end of the principal tentacle; the
whitish spot is a radiolarian. J, apical view. (Figs. 6B-J: © Bathybiologica A/S).
SCI. MAR., 70(3), September 2006, 363-379. ISSN: 0214-8358
NEW SPECIES OF PLANKTONIC CNIDARIANS FROM THE GULF OF MAINE • 371
is oblique and higher than it is wide, and although L.
lelouveteau somatocyst is very similar to L. quadriculata, the former species shows a marked velar ridge
and the ventro-basal margins of the hydroecium are
well rounded (Totton, 1965). Examination of three
anterior nectophores of L. lelouveteau collected in
Suruga Bay (Pagès, unpublished data) confirmed the
lack of horizontal ridges in this species. We prefer to
assign the new species to the genus Lensia based on
the presence of the five sets of vertical ridges. The
discovery of the eudoxid would keep it in the genus
Lensia or move it to Gilia.
Sub-Class ANTHOMEDUSAE Haeckel, 1879
Order FILIFERA Kühn, 1913
Family BOUGAINVILLIIDAE, Lütken, 1850
Bougainvilllia principis (Steenstrup, 1850)
(Fig. 5B-E)
Material: One large specimen collected in Georges Basin
(42º25.3’N, 67º31.3’W) between 290 m depth and the sea surface
on 25 September 2004. Bottom depth 297 m. Seawater temperature
on surface, 50 m and 270 m depth was 15.4, 4.7 and 8.2ºC respectively. Salinity at these depths was 32.0, 32.8 and 34. 9 psu respectively.
Description: Umbrella globular, 12 mm high and
11 mm wide, with homogenously thick mesoglea
that extends as “epaulettes” in the interradial segment below the subumbrella opening (Fig. 5B-E).
No peduncle. Manubrium cruciform, yellowish,
with long arms, high adradial stomach walls, narrow
perradial sides and deep interradial furrows that
occupy half of the subumbrella height. Radial canals
4, linear, wide. Ring canal as wide as radial ones.
Gonads flat, whitish, covering the adradial stomach
walls almost completely. Basal trunk of oral tentacles very short, branched six times, with a terminal
cnidocyst cluster that reaches the subumbrellar
opening when relaxed. Marginal bulbs ∧-shaped,
red-brown in colour, with about 40 short tentacles
per bulb arranged in a single row although this can
look like two rows due to bulb contraction; each tentacle with a rounded, black adaxial ocelli at its base.
Velum moderately broad.
Remarks: The specimen is a large adult according
to the umbrella size, shape and size of the manubrium, gonads, and number of tentacles. The description
fits quite well with the diagnosis for Bougainvillia
principis given by Kramp (1959), by Vanucci and
Rees (1961, Table VI) in their review of the genus
Bougainvillia, and by Edwards (1966, Table 4).
However, no figures in the literature illustrate a specimen similar to that reproduced in this study.
The ∧-shape of the marginal bulbs could
misidentify the species since another three species,
Bougainvillia macloviana Lesson, 1843, B. multitentaculata Foerster, 1923 and B. superciliaris (L.
Agassiz, 1849) (see Table VI in Vanucci and Rees,
1961) are known to have it. But these three species
have peduncle. However, we believe that tentacular
bulbs become ∧-shaped in large specimens due to
umbrella contraction. The re-examination of young
specimens of B. macloviana from the Magellan
channels (Pagès and Orejas, 1999) revealed that all
of them displayed linear shaped bulbs. Likewise
Figures 5 B-D show that the lower half of the
umbrella is not as wide as the upper one, which suggests that the living specimen contracted when the
photographs were taken.
Our specimen shows a short mesogleal space
above the stomach that could be interpreted as a
peduncle. But this space is not the case since the
radial canals are connected with the stomach base
without turning down and therefore without giving
rise to a peduncle. After preservation, this mesogeal
space became even more reduced. The adradial
walls of the stomach are rectangular shaped.
From the hydroid, Edwards (1966) reared
medusae up to 6.19 mm in diameter and 5.3 mm in
height, which was the first stage that he considered
fully adult. Both the number of tentacles per bulb
and the branching of oral tentacles in this specimen
were 19 and 8 respectively. The largest medusae collected in the Firth of Clyde waters measured up to
6.55 mm in diameter, had 23 tentacles per bulb and
oral tentacles branched 6 to 7 times.
To our knowledge the geographical position of
the specimen captured represents the most southern
and western record in the north Atlantic (Kramp,
1959; Shih et al., 1971; Cairns et al., 2002). Kramp
(1959) placed this species in a group of nine with
predominant northern-boreal or arctic-boreal distribution. A possible explanation for this occurrence is
the transport of the specimen from Greenland waters
and its advection into Georges Basin through the
Northeast Channel, which is supported by its large
size and mature stage. Kramp (1942) found several
specimens (2 to 11 mm wide) on the south-western
coast of Greenland. The hydrography of Georges
Bank and the adjacent Gulf of Maine is very complex and there are nine water masses within an area
of about 300 km in diameter (Flagg, 1987).
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372 • F. PAGÈS et al.
Bougainvilllia platygaster (Haeckel, 1879)
(Fig. 5F)
Material: One specimen collected in Oceanographer Canyon waters
(40º09.0’N 68º09.8’W) in the 200 to 100 m depth range on 16
September 2004.
Description: Umbrella globular, slightly cubic, 5
mm high and 5 mm wide, with homogenous thick
mesoglea that extends as “epaulettes” in the interand adradial segments below the subumbrellar
opening. No peduncle. Manubrium short, one-third
of subumbrellar height, with wide square base.
Radial canals 4, wide, linear. Ring canal narrow.
Gonads interradial, flat, yellowish. Basal trunk of
oral tentacles short, branched four times, with a terminal cnidocyst cluster. Marginal bulbs slightly
crescent, red-brown in colour, with about 10 long
tentacles per bulb, arranged in one row; each tentacle with a rounded, black adaxial ocelli at its base.
Velum moderately broad.
Remarks: Medusa buds were not observed on the
radial canals of the specimen but this absence was
not considered of taxonomic value by Bouillon
(1995, p. 227). He also noticed the lack of good
illustrations of the specific characters of this species
in the literature. In life, the manubrium is short (Fig.
5F), not as short as shown in drawings of preserved
specimens in which it looks flat (e.g. Pagès et al.,
1992, Fig. 2; Schuchert 1996, Fig. 16) but as it was
illustrated by Bouillon (1995, Fig. 1A). Cairns et al.
(2002) did not report this species in US and
Canadian waters.
Family JEANBOUILLONIIDAE fam. nov.
Diagnosis: as for the genus Jeanbouillonia.
Jeanbouillonia maserati gen. nov. et sp. nov.
(Figs. 5G-J, 6)
Material: One specimen collected in Georges Basin (42º25.3’N
67º31.3’W) between 290 m depth and the sea surface on 25
September 2004. Bottom depth 297 m. Holotype deposited in the
Smithsonian Institution.
Etimology: This genus was named after Prof.
Jean Bouillon, in recognition of his remarkable
work in the biology and systematics of hydrozoans
and for his friendship and support through many
years. The specific name maserati (name in apposition) is given for the trident-shaped main tentacle,
figuratively a nexus between the three families.
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FIG. 6. – Lateral view of the holotype of Jeanbouillonia maserati,
gen. nov. et sp. nov., umbrella 3 mm high.
Description: Umbrella globular, symmetrical,
without apical projection, 3 mm in diameter. The
subumbrellar cavity occupies the lower half of the
umbrella. About 1.5 mm of mesoglea above the subumbrella. Manubrium cruciform, orange, 1 mm
high, with perradial ridges and deep interradial furrows, filling almost all the subumbrellar cavity.
Mouth with four intangible lips that reach the subumbrellar opening. Radial canals 4, as narrow as the
ring canal. Gonads flat, yellowish, interradial,
extended over the whole manubrium except the perradial ridges. Gonads look like a tongue, wide in the
adradius and connected in the lower interradial half
of the manubrium, with a vertical furrow in the
upper half. Marginal bulbs 4, orange in colour, triangular, isosceles, with 7 to 8 cnidocysts on the top
corner, one with a tentacle and three without. One
cnidocysts line departs from each marginal bulb
upwards to mid-exumbrella. Tentacle 1.5 mm long,
trident-shaped due to three prominent orange terminal branches, with cnidocyst clubs arranged longitudinally on one side. The opposite marginal bulb has
no evident tentacle but a rounded knob with cnidocysts at its base; the two other marginal bulbs are
rudimentary, atentaculated.
NEW SPECIES OF PLANKTONIC CNIDARIANS FROM THE GULF OF MAINE • 373
Diagnosis of the genus Jeanbouillonia: Umbrella
globular. Exumbrellar cnidocyst track departing
upwards from each marginal bulb. Subumbrellar
cavity in the lower half of the umbrella. Manubrium
cruciform. Mouth with four simple lips. Radial
canals 4, linear. Gonads interradial, extended over
the whole manubrium except the perradial ridges.
Marginal bulbs 4. Only one tentacle, trident-shaped;
opposite marginal bulb with a knob; two remnant
bulbs rudimentary, atentaculated.
Remarks: Radiolarians were attached both to the
exumbrellar summit and at the base of the forked
distal end of the marginal tentacle but they were not
removed in order to avoid damaging the only specimen. However, the photographs of the living specimen (Fig. 5G-J) taken shortly after its capture disregard any interpretation of damage. The
Euphysora-type
morphological
appearance
(umbrella shaped plus arrangement, shape and
number of marginal tentacles; see Bouillon (1978)
for a species key) of this anthomedusan contrasts
strongly with the cruciform manubrium, the interradial gonads and the simple lips. These three characters neither match with the diagnosis of corymorphids nor with that for tubularids to which the
exumbrellar cnidocyst patch could relate. This
exumbrellar cnidocyst track is also characteristic of
zancleids (see Russell, 1953; Boero et al., 2000;
Bouillon et al., 2004) and a few pandeids. Zancleids
also share more characters with the new species,
namely the interradial gonads and the lateral position of cnidocysts in the tentacle, which nonetheless
are not developed in Jeanbouillonia as cnidophores
are in zancleids. The four simple lips of
Jeanbouillonia are rare but they have also been
described in the zancleid Halocoryne frasca Boero,
Bouillon and Gravili, 2000, whose tentacular bulbs
are also triangular and shelter cnidocysts.
In summary, the morphological features
described above are partially shared by at least
three families (corymorphids, tubularids and zancleids) but combined together do not fit any
known anthomedusan family, which suggests that
a new family needs to be established to accommodate this new genus and species in the hydrozoan
systematics. We believe that the effort of fitting
Jeanbouillonia maserati into any of the above
mentioned families, more than being an unconvincing adjustment would cause confusion in
hydrozoan systematics.
Subclass LEPTOMEDUSAE Haeckel, 1866
Order CONICA Broch, 1910
Family CAMPANULARIIDAE Johnston, 1837
Clytia hemisphaerica (Linnaeus, 1767)
Material: One specimen collected by dipnet in Georges Basin
(42°15.8’N, 67°29.1’W) on 13 September 2003. Many specimens
collected in surface waters off Gloucester Harbour (42°32.1’N
70°37.4’W) between 9 and 14 September 2004. Many more specimens collected at the top 10 m depth of Georges Basin waters
(42°26’N, 67°31.0’W) on 24 to 26 September 2004.
Description: Umbrella hemisphaerical, up to 15
mm in diameter. Gonads along the distal half of the
four radial canals. Tentacles 32, statocysts 2-3 with
one statolith between adjacent tentacles.
Remarks: Kramp (1959, p. 147) separated Clytia
(Phialidium) languida and C. hemisphaerica in the
identification key of the Phialidium species based
only on geographic distribution and stated “This
medusa (Clytia languida) may hardly be distinguished from P. hemisphaericum, ... their hydroids,
however, seem to be different”. Calder (1991, p. 5860) reviewed the Clytia species inhabiting northwestern Atlantic waters and indicated that both
species were probably the same. We concur with this
conclusion after examining our specimens. Cairns et
al. (2002) did not report C. languida but rather C.
hemisphaerica on the Atlantic and Pacific coasts of
USA and Canada.
Family ORCHISTOMIDAE Bouillon, 1984
Orchistoma collapsum (Mayer, 1900)
(Fig. 7A-C)
Material: One specimen collected in Oceanographic Canyon on 16
September 2004 in the 100 to 5 m depth range. Temperature 13.0 to
28 3ºC, salinity 34.9 to 36.5 psu.
Description: Umbrella wider (12 mm) than it is
high (9 mm), with a large, gelatinous gastric peduncle. The stomach has a broad H-shape base, with
low walls that meander along the peduncle forming
four substomachs separated by large ∩-shape spaces
(Fig. 7A). These substomachs branch dichotomiously giving rise to 8 substomachs with higher stomach
walls and crenulated lips, also separated by ∩-shape
spaces. The radial canals are linear, wide, greenish
and slightly jagged on the distal end. They depart
from the base of the 8 substomachs, arranged in
groups of 5,3,5,3,5,3,5 and 4 that sum 33 radial
canals in total. The short gonads are located in the
proximal part of the radial canals, very close to the
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374 • F. PAGÈS et al.
FIG. 7. – Orchistoma collapsum: A, subumbrellar view, umbrella diameter 12 mm. B and C, abaxial view of the umbrella margin showing
tentacle bulbs, secondary tentacles and adaxial ocelli. Dichotomia cannoides: D, subumbrellar view, umbrella diameter 3.5 mm. E, apico-lateral view, umbrella 5 mm high. Leuckartiara octona: F, lateral view, umbrella 15 mm high. Cytaeis spp.: G, lateral view of umbrella 3 mm
high. Crossota rufobrunnea: H, subumbrellar view, 11 mm in diameter. Nausithoe punctata: I, subumbrellar view, 10 mm in diameter; note
the specific circular ocelli in the rhopalia. Beroe sp.: J, oral-aboral length 22 mm, with a whitish prey in the pharynx. (Fig. 7A-J:
© Bathybiologica A/S).
basal, external perimeter of the stomach. The eggs
measure 0.2 mm in diameter and are arranged in two
rows along the gonads. The marginal rim has 33
vesicles, each with endodermal dark spots. The primary marginal tentacles are cut at their base; the
secondary ones, 4 to 12 between adjacent vesicles,
are short, 0.3 mm in length. Ocelli adaxial, dark,
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rounded, located in the ring canal without corresponding to the tentacles (Fig. 7 B-C).
Remarks: Bouillon (1984a, p.15) created the family Orchistomidae and included a diagnosis in an
attempt to make a new classification for ThecataeLeptomedusae. The new family was, however, justi-
NEW SPECIES OF PLANKTONIC CNIDARIANS FROM THE GULF OF MAINE • 375
fied in the following paper (Bouillon, 1984b, p. 88),
in regrouping several medusae species formerly
placed in the Dipleurosomatidae as Dipleurosoma
collapsum (Mayer 1900), or in the Laodiceidae as
Orchistoma agariciforme Keller, 1884 and O. pileus
(Lesson, 1843) among others. Bouillon (1984b)
agreed with Kramp (1959, p.132) that some
Dipleurosoma species probably did not belong to this
genus because the rudimentary tentacles were not
laodiceid cordyli. However, illustrations of the anatomy of the tentacles were not provided. The overall
good shape of our specimen allowed a microscopical
observation of the umbrella margin, which confirmed
that the tentacles were rudimentary, without any
resemblance to laodiceid cordyli (Fig. 7B-C).
Family DIPLEUROSOMIDAE Russell, 1953
Dichotomia cannoides Brooks, 1903
(Fig. 7D-E)
Material: One specimen collected in Oceanographic Canyon
(40º09.0’N, 68º09.8’W) between 100 and 5 m depth on 16 September
2004. Temperature 13.0 to 28.3ºC, salinity 34.9 to 36.5 psu.
Additional material examined: More than 50 specimens collected in
Papua New Guinea waters by Prof. Jean Bouillon and deposited in
the biological collections of the Institut Royal des Sciences
Naturelles de Belgique.
Description: Umbrella bell-shape, 5 mm high,
3.5 mm wide. Rounded apical projection with
mesoglea moderately thick. Radial canals 29, almost
as wide as interradial spaces, orange-brown in
colour, derived from a central H-shape canal system
in the subumbrella top, whose 4 arms bifurcate three
times, although not always regularly. Gonads folded
and mainly located along the three bifurcations and
the proximal part of the definitive radial canals.
Tentacles solid, about 60, all of similar structure and
length, densely crowded, without keeping a numeric correspondence with the radial canals.
Manubrium not found.
Remarks: The upward orientation of the tentacles
(Brooks, 1903; Kramp, 1959) could be a preservation artifact as it was not observed in the living specimen (Fig. 7 D-E) or in the specimens deposited in
Brussels. As Kramp (1959) wrote, and contrary to
Brooks’s description, we observed only one type of
solid tentacle, which was variable in length due to
contraction as the shorter tentacles have more
rounded distal ends. However, the specimens
deposited in Brussels have tentacles of two sizes like
those originally illustrated.
As there was no proper manubrium in our specimen and the one figured by Brooks was odd, large
and simple (1903, plate 1 Fig. 1), which was also
copied and poorly described by Kramp (1959) in
spite of having found 14 specimens 3 to 8 mm in
diameter, we decided to examine several specimens
collected by Prof. Jean Bouillon in waters of Papua
New Guinea. These specimens range between 2 and
5 mm in height and none of them has a tubular
manubrium with simple mouth. However, all of them
have a polygonal manubrium that occupies almost
half the subumbrella in the largest individuals, with
the same number of lips and branches before the last
bifurcation. This relation led us to believe that the
manubrium of our specimen could have about 14
lips. This species shows a wide geographical distribution with few records: Bahamas Islands (Brooks,
1903), Virgin Islands and Delaware Bay (Kramp,
1959), South China Sea (Xu and Zhang, 1978) and
Papua New Guinea (Bouillon, 1984).
Family TIARANNIDAE Russell, 1940
Modeeria rotunda (Quoy and Gaimard, 1827)
(Fig. 8A-C)
Material: One specimen collected in Oceanographer Canyon
(40º09.5’N, 67º57.4’W) between 494 and 378 m depth on 7
September 2002. Seawater temperature 5.7 to 6.4ºC, salinity 34.98
to 35.02 psu.
Description: Umbrella globular, 24 mm high and
26 mm wide in the base. Mesoglea very thick, 6 mm
from stomach base to umbrella summit.
Subumbrellar cavity pyriform (Fig. 8A).
Manubrium red, cruciform, looking like a pyramid
of 8 sides in apical view, the wider sides perradial,
the narrower interradial, reaching mid-subumbrella.
The manubrium extends as four perradial pouches
attached to the subumbrella (Fig. 8B) (Bouillon,
1985, p. 144). Radial canals 4, thin, that depart from
the lower third of the manubrium, below the lower
end of the gonads (Fig. 8B), and bend inwards following four, ovoid, mesogleal convexities that can
look bilobate (Fig. 8C). The radial canals run downwards to the ring canal but one of them bifurcates
beneath the mesogleal convexity (Fig. 8A) and five
join the ring canal. Mouth large with four lips slightly crenulated. Subumbrellar opening slightly square.
Tentacles 32, whitish, with slight lateral crenulations. Tentacular bulbs with red, rounded endoderm.
Gonads adradial, forming horizontal folds on the
manubrium and perradial gastric pouches, connected apically in the interradii and filling all the
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376 • F. PAGÈS et al.
FIG. 8. – Modeeria rotunda, 24 mm umbrella height: A, lateral
view. Note the pyriform subumbrella with the oval-shape perradial
convexity. B, detail of the gonads and the perradial convexity. C,
subumbrellar view of the manubrium surrounded by the four
perradial convexities.
manubrial subumbrella. Cordyli-like structures 1 to
3, usually 2, between marginal tentacles.
Remarks: Modeeria (Tiaranna) rotunda descriptions show remarkable differences in the growth
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stage and size when comparing the text and figures
from several sources in the literature (Ranson, 1936;
Kramp, 1959; Edwards, 1963, 1973). For instance,
the pyriform subumbrella and the four conspicuous
mesogleal convexities characteristic of large specimens are rarely mentioned. To our knowledge,
Hartlaub (1917, Fig. 341 as Rotundula brochii) provided the first and only drawing of the pyriform subumbrella from a specimen 25 to 30 mm in diameter.
A morphology which was also shown in the photograph of a large specimen published by Wrobel and
Mills (1998, p. 34). The mesogleal convexities were
described by Ranson (1936, p. 68, Pl. 1, Figs. 10-11)
as mesogleal spurs (éperons). It is interesting to note
that the specimen figured (Ranson 1936, Pl. 1, Fig.
11) had 5 radial canals and 5 mesogleal convexities.
The mesogleal convexities were photographed by
Brinckmann-Voss and Arai (1998, Fig. 12C) in a
specimen 20 mm wide and 17 mm high. They
argued that the inwards bending of the radial canals
and the connection to the manubrium formed a subumbrellar pocket. However, Edwards had used the
last term previously (1973, Fig. 2 A, C-D) for designating the interradial spaces above the pyramidal
manubrium in a specimen 8.46 mm in diameter and
6.37 mm high. The use of the term “subumbrellar
pocket” could cause some confusion in interpreting
explanations from both papers. For example, neither
of these spaces exists in our specimen due to the
large size of the gonads. Brinckmann-Voss (pers.
comm. on 11 March 2005) believes that “subumbrellar pocket” was a poor expression and that
Edwards could not refer to the same space because
of the young age of his specimens.
The position of the gonads varies depending on
each author. Edwards (1973, p. 577) wrote “ Gonads
have begun to form as sinuous folds in the walls of
the stomach immediately adjacent to its perradial
attachments to the subumbrella...”. Bouillon partially concurred (1985, p. 145) in considering that they
are located over the interradial surface of the
manubrium as well as over the perradial lobes.
Ranson (1936) and Brinckmann-Voss and Arai
(1998) placed them as adradial. We believe that the
three explanations are correct but adradial is the
most exact location because perradial is the attachment of the stomach pouches and the interradii are
free of gonads except in the apical part of the
manubrium (Fig. 8). Cairns et al. (2002) reported
this species only on the Pacific coast of USA and
Canada.
NEW SPECIES OF PLANKTONIC CNIDARIANS FROM THE GULF OF MAINE • 377
FIG. 9. – Pleurobrachia pileus: A, specimen 20 mm in oral-aboral length with 6 cestodes marked by arrows. B, two cestodes located in the
canal of the infundibulum, the longest is 2.4 mm. C, detail of the four suckers of the cestode head.
Phylum CTENOPHORA Eschscholtz, 1829,
Class TENTACULATA Eschscholtz, 1825
Order CYDIPPIDA Gegenbaur, 1856
Family PLEUROBRACHIDAE Chun, 1880
Pleurobrachia pileus (O.F. Müller, 1776)
(Fig. 9A-C)
of cestodes. However, cestodes are difficult to identify and a life cycle involving coelenterates has not
been established yet (Lauckner, 1980; Purcell and
Arai, 2001). Specimens of Pleurobrachia were not
collected in 2002 but several were found in Georges
Basin in 2004.
Material and remarks: In 2003, many specimens
(1.5 to 2.0 cm in oral-aboral length) were collected in
the top 35 m of Georges Basin waters. Some of them
were parasitized by cestodes (1.6-2.4 mm in length)
ranging from 1 to 6 per ctenophore (Fig. 9 A-C). The
cestodes were located in the canal of the infundibulum, the pharynx or the tentacle sheath of the
ctenophores. This association was reported off
Woods Hole by van Cleave (1927) who found four
immature cestodes (1.5 mm in length, preliminarily
identified as Scolex polymorphus by Edwin Linton)
in three collected specimens of Pleurobrachia pileus,
but no further observations were noted. We could not
identify the cestode species. It is known that the
names Scolex polymorphus and S. pleuronectis, both
of which have already been found in ctenophores,
encompass several cestode genera (Stunkard, 1977,
p. 402 for an updating). Ctenophores are sometimes
the main prey of fishes (e.g. Mianzan et al., 1996)
and play a role as intermediate hosts in the life-cycle
Class NUDA Chun, 1879
Order BEROIDA Eschscholtz, 1829
Family BEROIDAE Eschscholtz, 1829?
Beroe sp. (Figs. 7J and 10)
Material: Few specimens were collected in the first cruise
(Oceanographer Canyon, 5 September 2002, 778-680 m, 1 specimen; 22 September 2002, 680-583 m depth, 1 specimen; 679 m
depth, 2 specimens. Hydrographer Canyon, 17 September 2002,
600 to 500 m depth, 4 specimens). In 2003, it was common and
widely distributed in the basins (Wilkinson, top 110 m depth;
Georges, top 210 m depth) with some individuals reaching 21 cm in
oral-aboral length. In 2004, it was often observed in surface waters
and several specimens were captured by the Meternet hauls in
Wilkinson and Georges basins but not in the canyons.
Remarks: The specimens have not been identified to species level following the recommendations of a genus review stated by Mills et al. (1996).
The Meternet haul that collected the two new
anthomedusae species also captured three small
transparent Beroe and a large pinkish one. The large
specimen (8 cm in oral-aboral length) was paraSCI. MAR., 70(3), September 2006, 363-379. ISSN: 0214-8358
378 • F. PAGÈS et al.
FIG. 10. – Beroe sp. with an amphipod Hyperia galba (identified by Rebeca Gasca) preying on it. The oral side was wrenched by a
molecular parasite.
sitized by a large, brown specimen of the hyperid
amphipod Hyperia galba that preyed upon the surface tissues through which it had made a hole (1 cm
in diameter, Fig. 10). The Beroe was placed in a
bowl and the hyperid remained attached to it,
devouring the hole edges.
CONCLUSIONS
This study reveals that the gelatinous fauna in the
Gulf of Maine comprise several species about which
little is known or which have not yet been described.
Siphonophores appear to have a high species richness (33 of the 160 species described to date in
world oceans), while hydroidomedusae is proportionally the less rich taxon (26 of approx. 842
species described worldwide, see Bouillon and
Boero, 2000). The scyphomedusae collected were
almost the same species reported by Larson et al.
(1988). The number of mollusc and salp species was
low. A comparison with the gelatinous species
occurring in the Gulf of Maine and discussed by
Bigelow (1926) shows some species that occur in
both studies (Nanomia cara, Physophora hydrostatica, Melicertum octocostatum, Clytia hemisphaerica, Phacellophora camstchatica, Periphylla periphylla, Beroe sp., Pleurobrachia pileus) but most of
the rest of the species do not.
These results encourage the exploration of the
gelatinous plankton populations in the Gulf of
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Maine and the rest of the world’s oceans because
many species and biological relationships remain
unknown.
ACKNOWLEDGEMENTS
The scientific results of this study and the publication have been funded by the Belgian Fonds Léopold
III pour l’Exploration et la Conservation de la
Nature. We thank the captains and crew of the Seward
Johnson vessels for assisting with all aspects of animal collection. The Institut Royal des Sciences
Naturelles de Belgique is acknowledged for the loan
of biological material deposited there. The Japanese
Society for Promotion of Science supported FP in a
part of this study. This research was supported by
grants to MY from the National Science Foundation
(OCE 0002493) and the Harbor Branch
Oceanographic Institution. This study has been conducted in the frame of the Census of Marine
Zooplankton project. We thank two anonymous referees by their useful comments. Harbor Branch
Oceanographic Institution Contribution No. 1637.
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Scient. ed.: P. Abelló
Received April 1, 2005. Accepted March 13, 2006.
Published online July 5, 2006.
SCI. MAR., 70(3), September 2006, 363-379. ISSN: 0214-8358